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          细胞质基质与内膜系统
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            <p>细胞生物学 第7章 细胞质基质与内膜系统</p>
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<a id="more"></a>

<hr>
<p>细胞内被膜区分为3 类结构：</p>
<ul>
<li>细胞质基质</li>
<li>内膜系统：是指在结构、功能乃至发生上相互关联、由单层膜包被的细胞器或细胞结构。主要包括内质网、高尔基体、溶酶体、胞内体和分泌泡等。</li>
<li>其他由膜包被的细胞器：诸如线粒体、叶绿体、过氧物酶体和细胞核。</li>
</ul>
<h2 id="细胞质基质及其功能"><a href="#细胞质基质及其功能" class="headerlink" title="细胞质基质及其功能"></a>细胞质基质及其功能</h2><p>细胞质基质：在真核细胞的细胞质中，除去可分辨的细胞器以外的胶状物质，占据着细胞膜内、细胞核外的细胞内空间，称细胞质基质。</p>
<blockquote>
<p>细胞质基质的主要成分包括约占总体积70％ 的水和溶于其中的离子以及以可溶性蛋白质为主的大分子， 其体积占细胞总体积50％以上。</p>
</blockquote>
<h3 id="细胞质基质的含义"><a href="#细胞质基质的含义" class="headerlink" title="细胞质基质的含义"></a>细胞质基质的含义</h3><h3 id="细胞质基质的功能"><a href="#细胞质基质的功能" class="headerlink" title="细胞质基质的功能"></a>细胞质基质的功能</h3><p>目前了解最多的是许多中间代谢过程都发生在细胞质基质中， 如糖酵解过程、磷酸戊糖途径、糖醋酸途径、糖原的合成与部分分解过程等。</p>
<p>细胞质基质的首要功能是为某些蛋白质合成和脂肪酸合成提供场所。</p>
<p>细胞质基质第二方面的功能是与细胞质骨架相关的。</p>
<p>细胞质基质第三方面的功能是与细胞膜相关的。</p>
<p>细胞质基质第四方面功能， 是与蛋白质的修饰和选择性降解等方面有关。</p>
<p>蛋白质的修饰；控制蛋白质的寿命；降解变性和错误折叠的蛋白质；帮助变性或错误折叠的蛋白质重新折叠， 形成正确的分子构象</p>
<h2 id="细胞内膜系统及其功能"><a href="#细胞内膜系统及其功能" class="headerlink" title="细胞内膜系统及其功能"></a>细胞内膜系统及其功能</h2><h3 id="细胞内膜系统"><a href="#细胞内膜系统" class="headerlink" title="细胞内膜系统"></a>细胞内膜系统</h3><p>细胞内膜系统：是指在结构、功能乃至发生上相互关联、由单层膜包被的细胞器或细胞结构， 主要包括内质网、高尔基体、溶酶体、胞内体和分泌泡等。</p>
<p>主要研究技术：</p>
<ul>
<li>电镜技术：揭示超微结构</li>
<li>放射自显影技术：用于功能定位研究</li>
<li>差速离心技术：组分分离及功能分析</li>
<li>荧光显微技术：大分子（GFP）标记</li>
<li>突变体分析技术：膜泡运输和遗传基础</li>
</ul>
<h3 id="内质网"><a href="#内质网" class="headerlink" title="内质网"></a>内质网</h3><p>内质网（ER）是真核细胞中最普遍、最多变、适应性最强的细胞器。</p>
<p>内质网由封闭的<strong>管状</strong>或<strong>扁平囊状膜</strong>系统及其包被的腔形成互相沟通的三维网络结构。</p>
<p>内质网通常占细胞膜系统的一半左右，体积占细胞总体积的10%以上。</p>
<p>在不同类型的细胞中，内质网的数量、类型与形态差异很大， 同一细胞在不同发育阶段和不同的生理状态下， 内质网的结构与功能也随之显著变化。在细胞周期不同阶段， 内质网也发生很大变化。细胞分裂时， 内质网要经历解体与重建的过程。</p>
<hr>
<p>微粒体：细胞匀浆和超速离心过程中， 由<strong>破碎的内质网</strong>形成的近似球形的囊泡结构， 它包含内质网膜与核糖体两种基本组分。</p>
<blockquote>
<p>但在体外实验中， 微粒体仍具有蛋白质合成、蛋白质的糖基化和脂质合成等内质网的基本功能。因此在生化与功能研究中， 常常把微粒体和内质网等同看待。</p>
</blockquote>
<hr>
<p>通过微粒体研究，内质网的主要化学成份是蛋白质和脂类。<br>蛋白质（60-70%）：代谢酶系</p>
<ul>
<li>NADPH细胞色素P450还原酶</li>
<li>NADPH-细胞色素b5还原酶</li>
<li>细胞色素b5、细胞色素P450、脂肪酸CoA连接酶等</li>
</ul>
<p>标志酶：葡萄糖-6-磷酸酶</p>
<p>脂类（30-40%）：以磷脂酰胆碱为主，比细胞膜丰富，鞘磷脂少</p>
<hr>
<p>内质网是细胞内<strong>蛋白质</strong>与<strong>脂质</strong>合成的基地，几乎全部的脂质和多种重要的蛋白质都是在内质网上合成的。是行使多种重要功能的复杂结构体系</p>
<h4 id="基本类型"><a href="#基本类型" class="headerlink" title="基本类型"></a>基本类型</h4><p>糙面内质网（rER ）和光面内质网（sER）</p>
<h5 id="糙面内质网"><a href="#糙面内质网" class="headerlink" title="糙面内质网"></a>糙面内质网</h5><p>膜表面附着核糖体，膜上有核糖体连接蛋白；形态多为板层状排列的扁囊；网腔内含低电子或中等电子密度的物质；多分布在分泌活动旺盛或分化较完善的细胞内</p>
<blockquote>
<p>划分细胞分化程度的一个标准</p>
</blockquote>
<hr>
<p>糙面内质网多呈扁囊状， 排列较为整齐， 因其膜表面附有大量的核糖体而命名</p>
<p>多呈扁囊状， 排列较为整齐， 是内质网与核糖体共同形成的复合机能结构；</p>
<p>它是分泌性蛋白和多种膜蛋白合成场所；</p>
<hr>
<p>【功能一】蛋白质的合成</p>
<p>在核糖体上合成，并起始于细胞质基质，有些蛋白质在合成开始不久后便转到内质网上合成。</p>
<p>三种类型蛋白：</p>
<p>分泌蛋白，如：胰腺细胞分泌的消化酶；浆细胞分泌的抗体；小肠杯状细胞分泌的粘蛋白；内分泌腺细胞分泌的多肽激素；胞外基质成分等。</p>
<p>膜的整合蛋白（膜蛋白）：细胞膜上的膜蛋白、ER、高尔基体、溶酶体膜上的膜蛋白都由rER合成，然后定向转输到ER、高尔基体、溶酶体、质膜上。</p>
<p>构成细胞器中的可溶性驻留蛋白：溶酶体和植物液泡中的酸性水解酶；内质网、高尔基体、胞内体中固有蛋白</p>
<hr>
<p>【功能二】蛋白质的修饰与加工</p>
<p>进入内质网的蛋白质发生的化学修饰主要有糖基化、羟基化、酰基化、二硫键形成等。<br><strong>最主要的是糖基化</strong>，几乎所有内质网上合成的蛋白质最终被糖基化。</p>
<p><img src="../../../../../Pictures/blogimg/image-20200403220338303.png" alt="image-20200403220338303"></p>
<p><img src="../../../../../Pictures/blogimg/image-20200403220507464.png" alt="image-20200403220507464"></p>
<p>N-连接的糖基化，寡糖基转移到天冬酰胺残基上，与之连接的糖为N-乙酰葡糖胺，均发生在内质网中</p>
<p>O-连接的糖基化，寡糖基转移到丝氨酸、苏氨酸残基上，与之连接的糖为N-乙酰半乳糖胺，主要发生在高尔基体中</p>
<p><img src="../../../../../Pictures/blogimg/image-20200403220630852.png" alt="image-20200403220630852"></p>
<hr>
<p>【功能三】新生多肽的折叠与装配</p>
<p>肽链的合成仅需几十秒至几分钟，在内质网停留的时间往往长达几十分钟</p>
<p>内质网腔二硫键形成</p>
<p>附着在内质网膜腔面上的蛋白二硫键异构酶（PDI）可以切断错误的二硫键，形成自由能最低的蛋白质构象，从而帮助新合成的蛋白质重新形成二硫键并产生正确折叠的构象</p>
<p>内质网中的一种结合免疫球蛋白蛋白（Bip）<br>是属于HSP70家族的<strong>分子伴侣</strong>，在内质网中有2个作用：</p>
<ul>
<li>一是Bip同进入内质网的未折叠蛋白质的疏水氨基酸结合，防止多肽链不正确地折叠和聚合，或者识别错误折叠的蛋白质或未装配好的蛋白质亚单位，并促进它们重新折叠与装配；</li>
<li>二是防止新合成的蛋白质在转运过程中变性或断裂</li>
</ul>
<p>蛋白二硫鍵异构酶（PDI）和结合蛋白（Bip,HSP70）等蛋白质都具有<strong>4肽驻留信号</strong>（KDEL或HDEL）以保证它们滞留在内质网中，并维持很高的浓度（如同监狱）</p>
<p>内质网是蛋白质分泌转运途径中行使质量监控的重要场所。不能正确折叠的畸形肽链或未组装成寡聚体的蛋白质亚基，不论在內质网膜上还是在内质网腔中，一般都不能进入高尔基体，并通过Sec61p复合体从内质网腔转至细胞质基质，进而通过<strong>泛素依赖性降解途径</strong>被蛋白酶体所降解。</p>
<h5 id="光面内质网"><a href="#光面内质网" class="headerlink" title="光面内质网"></a>光面内质网</h5><p>表面没有附着核糖体的内质网称光面内质网， 光面内质网常为分支管状（泡状）， 形成较为复杂的立体结构。</p>
<p>光面内质网是脂质合成的重要场所</p>
<p>脂质的合成与转运</p>
<hr>
<p>【功能一】sER是脂质合成的重要场所</p>
<p>膜脂甾类激素</p>
<p>内质网合成构成细胞所需要的包括磷脂、固醇类在内的<strong>几乎全部的膜脂</strong>其中最主要的磷脂是磷脂酰胆碱</p>
<p>合成磷脂所需要的3种酶活性部位都定位在内质网膜的细胞质基质侧</p>
<p><img src="../../../../../Pictures/blogimg/%E5%8D%B5%E7%A3%B7%E8%84%82%E5%9C%A8%E5%86%85%E8%B4%A8%E7%BD%91%E8%86%9C%E4%B8%8A%E5%90%88%E6%88%90%E8%BF%87%E7%A8%8B%E7%9A%84%E7%A4%BA%E6%84%8F%E5%9C%88.png" alt="卵磷脂在内质网膜上合成过程的示意圈"></p>
<p><img src="../../../../../Pictures/blogimg/image-20200403214907295.png" alt="image-20200403214907295"></p>
<p><img src="../../../../../Pictures/blogimg/%E7%A3%B7%E8%84%82%E8%BD%AC%E4%BD%8D%E8%9B%8B%E7%99%BD%E6%88%96%E7%A7%B0%E8%BD%AC%E4%BD%8D%E9%85%B6.png" alt="磷脂转位蛋白或称转位酶"></p>
<hr>
<p>磷脂由内质网向其他膜的转运主要有3种方式</p>
<ul>
<li><p>以出芽的方式转运到高尔基体、溶酶体和细胞质膜上。</p>
</li>
<li><p>凭借一种水溶性的载体蛋白—磷脂交换蛋白（PEP）将內质网上合成的脂转运到线粒体、叶绿体和过氧化物酶体的膜中</p>
</li>
<li><p>通过膜嵌入蛋白所介导的直接接触转运膜脂。</p>
</li>
</ul>
<hr>
<p>【功能二】类固醇激素的合成</p>
<p>分泌类固醇激素的细胞如肾上腺细胞、睾丸间质细胞和黄体细胞都有丰富的光面内质网，并在光面内质网上含有合成胆固醇和将胆固醇转化为激素的全套酶系;</p>
<p>所以光面内质网能够合成胆固醇，然后将胆固醇氧化、还原、水解进一步转变成各种类固醇激素。</p>
<hr>
<p>【功能三】解毒作用</p>
<p>肝细胞中含有丰富的sER，它是合成外输性脂蛋白颗粒的基地。这些光面内质网含有一些酶（如混合功能氧化酶细胞色素P450），介导氧化、还原和水解反应，使有毒物质由脂溶性转变成水溶性而被排出体外，此过程称为肝细胞的解毒作用。</p>
<hr>
<p>【功能四】糖原代谢</p>
<p>sER中的葡萄糖-6-磷酸酶将葡萄糖-6-磷酸水解生成葡萄糖和无机磷，释放游离的葡萄糖进入血液，供细胞之用。</p>
<hr>
<p>【功能五】贮积钙离子（钙库）</p>
<p>肌质网：心肌和骨骼肌细胞中的一种特殊的内质网，储存和释放Ca2+，参与肌肉收缩活动。</p>
<h4 id="内质网的功能"><a href="#内质网的功能" class="headerlink" title="内质网的功能"></a>内质网的功能</h4><p>光面内质网是脂质合成的重要场所</p>
<h3 id="高尔基体"><a href="#高尔基体" class="headerlink" title="高尔基体"></a>高尔基体</h3><h4 id="高尔基体的功能"><a href="#高尔基体的功能" class="headerlink" title="高尔基体的功能"></a>高尔基体的功能</h4><h3 id="溶酶体"><a href="#溶酶体" class="headerlink" title="溶酶体"></a>溶酶体</h3><h4 id="溶酶体的形态结构与类型"><a href="#溶酶体的形态结构与类型" class="headerlink" title="溶酶体的形态结构与类型"></a>溶酶体的形态结构与类型</h4><p>【分类】根据溶酶体处于完成其生理功能的不同阶段， 大致可分为初级溶酶体、次级溶酶体和残质体。</p>
<h4 id="溶酶体的功能"><a href="#溶酶体的功能" class="headerlink" title="溶酶体的功能"></a>溶酶体的功能</h4><ul>
<li><p>清除无用的生物大分子、衰老的细胞器及衰老损伤和死亡的细胞</p>
</li>
<li><p>防御功能</p>
</li>
<li><p>作为细胞内的消化“器官”为细胞提供营养</p>
</li>
<li><p>在分泌腺细胞中，溶酶体常常摄入分泌颗粒，参与分泌过程的调节</p>
</li>
<li><p>某些特定细胞程序性死亡，死亡后的细胞被周围吞噬细胞溶酶体消化清除。</p>
</li>
<li><p>参与受精过程中的顶体反应</p>
</li>
</ul>
<h4 id="溶酶体的发生"><a href="#溶酶体的发生" class="headerlink" title="溶酶体的发生"></a>溶酶体的发生</h4><h3 id="过氧化物酶体"><a href="#过氧化物酶体" class="headerlink" title="过氧化物酶体"></a>过氧化物酶体</h3>
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          <div class="post-toc motion-element"><ol class="nav"><li class="nav-item nav-level-2"><a class="nav-link" href="#细胞质基质及其功能"><span class="nav-number">1.</span> <span class="nav-text">细胞质基质及其功能</span></a><ol class="nav-child"><li class="nav-item nav-level-3"><a class="nav-link" href="#细胞质基质的含义"><span class="nav-number">1.1.</span> <span class="nav-text">细胞质基质的含义</span></a></li><li class="nav-item nav-level-3"><a class="nav-link" href="#细胞质基质的功能"><span class="nav-number">1.2.</span> <span class="nav-text">细胞质基质的功能</span></a></li></ol></li><li class="nav-item nav-level-2"><a class="nav-link" href="#细胞内膜系统及其功能"><span class="nav-number">2.</span> <span class="nav-text">细胞内膜系统及其功能</span></a><ol class="nav-child"><li class="nav-item nav-level-3"><a class="nav-link" href="#细胞内膜系统"><span class="nav-number">2.1.</span> <span class="nav-text">细胞内膜系统</span></a></li><li class="nav-item nav-level-3"><a class="nav-link" href="#内质网"><span class="nav-number">2.2.</span> <span class="nav-text">内质网</span></a><ol class="nav-child"><li class="nav-item nav-level-4"><a class="nav-link" href="#基本类型"><span class="nav-number">2.2.1.</span> <span class="nav-text">基本类型</span></a><ol class="nav-child"><li class="nav-item nav-level-5"><a class="nav-link" href="#糙面内质网"><span class="nav-number">2.2.1.1.</span> <span class="nav-text">糙面内质网</span></a></li><li class="nav-item nav-level-5"><a class="nav-link" href="#光面内质网"><span class="nav-number">2.2.1.2.</span> <span class="nav-text">光面内质网</span></a></li></ol></li><li class="nav-item nav-level-4"><a class="nav-link" href="#内质网的功能"><span class="nav-number">2.2.2.</span> <span class="nav-text">内质网的功能</span></a></li></ol></li><li class="nav-item nav-level-3"><a class="nav-link" href="#高尔基体"><span class="nav-number">2.3.</span> <span class="nav-text">高尔基体</span></a><ol class="nav-child"><li class="nav-item nav-level-4"><a class="nav-link" href="#高尔基体的功能"><span class="nav-number">2.3.1.</span> <span class="nav-text">高尔基体的功能</span></a></li></ol></li><li class="nav-item nav-level-3"><a class="nav-link" href="#溶酶体"><span class="nav-number">2.4.</span> <span class="nav-text">溶酶体</span></a><ol class="nav-child"><li class="nav-item nav-level-4"><a class="nav-link" href="#溶酶体的形态结构与类型"><span class="nav-number">2.4.1.</span> <span class="nav-text">溶酶体的形态结构与类型</span></a></li><li class="nav-item nav-level-4"><a class="nav-link" href="#溶酶体的功能"><span class="nav-number">2.4.2.</span> <span class="nav-text">溶酶体的功能</span></a></li><li class="nav-item nav-level-4"><a class="nav-link" href="#溶酶体的发生"><span class="nav-number">2.4.3.</span> <span class="nav-text">溶酶体的发生</span></a></li></ol></li><li class="nav-item nav-level-3"><a class="nav-link" href="#过氧化物酶体"><span class="nav-number">2.5.</span> <span class="nav-text">过氧化物酶体</span></a></li></ol></li></ol></div>
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